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Significance Forests are experiencing growing risks of drought-induced mortality in a warming world. Yet, ecosystem dynamics following drought mortality remain unknown, representing a major limitation to our understanding of the ecological consequences of climate change. We provide an emerging picture of postdrought ecological trajectories based on field indicators of forest dynamics. Replacement patterns following mortality indicate limited short-term persistence of predrought dominant tree species, highlighting the potential for major ecosystem reorganization in the coming decades. The great variability of the observed dynamics within and among species reinforces the primary influence of drought characteristics and ecosystem legacies, modulated by land use, management, and past disturbances, on ongoing drought-related species turnover and their potential implications for future forest biodiversity and ecosystem services., Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Recent, widespread spruce beetle (Dendroctonus rufipennis) outbreaks have driven extensive tree mortality across western North America. Post-disturbance forest management often includes salvage logging to capture economic value of dead timber, reduce fire hazard, and meet other social or ecological objectives. Little is known about effects of salvage logging on surface fuel loads or plant understory communities in Engelmann spruce (Picea engelmannii)-dominated forests. We sampled fine and coarse woody debris, ground cover, and plant species composition along transects in spruce-beetle impacted stands in southwestern Colorado, USA. Twenty stands had been subject to clearcut 1–2 years prior to sampling; 32 stands were unlogged controls. Salvage logged stands exhibited altered surface fuels, ground cover, plant species cover, and community composition. Salvage increased 1-, 10-, and 100-hr fuels, and cover by bare ground and woody debris; cover by litter and cryptogams was reduced. Understory plant cover was reduced in salvaged stands, primarily due to losses of shrub cover. We found no difference in species diversity or richness between treated stands and controls. Salvage logging also drove shifts in plant community composition. Mean cover by non-native species was low, and not different, between control and salvage stands. Our study characterizes short-term effects that will undoubtedly change substantially over longer periods, in particular due to anticipated tree seedling growth and movement of standing dead wood from the canopy to the surface. In the near term, abundant fine woody surface fuels at salvage sites could influence the likelihood and rate of spread of surface fires, though over time we expect surface fuels in untreated stands to increase to comparable or greater levels than in salvaged stands. Differences in vascular plant cover and composition imparted by salvage harvests is also expected to change over time, though whether treated and untreated communities diverge or converge is not known. As such, we recommend that salvage harvest effects be monitored over extended time frames in order to detect longer-term trends.

Fire regimes in many dry forests of western North America are substantially different from historical conditions, and there is concern about the ability of these forests to recover following severe wildfire. Fire refugia, unburned or low-severity burned patches where trees survived fire, may serve as essential propagule sources that enable forest regeneration. To quantify the influence of fire refugia spatial pattern and other biophysical factors on the process of post-fire tree regeneration; in particular examining both the proximity and density of surrounding refugia to characterize the landscape of refugial seed sources. We surveyed regeneration at 135 sites in stand-replacement patches across a gradient of fire refugia density in eastern Oregon, USA. We characterized the influence of refugial seed source pattern and other environmental factors on the abundance of regenerating seedlings, and examined the relationship between post-fire climate and the temporal pattern of ponderosa pine seedling establishment. Tree seedlings were present in 83% of plots 12–17 years post-fire, and densities varied substantially (0–67800 stems ha−1, median = 1100). Variation in seedling abundance was driven by the spatial patterns of refugial seed sources. Despite widespread post-fire shrub cover, high-severity burned forests have not undergone a persistent type conversion to shrublands. Ponderosa pine seedling establishment peaked 5–11 years after fire and was not closely associated with post-fire climate. Fire refugia and the seed sources they contain fostered tree regeneration in severely burned patches. Management practices that reduce refugia within post-fire landscapes may negatively influence essential forest recovery processes.

Author(s): Krawchuk, Meg A; Meigs, Garrett W; Cartwright, Jennifer M; Coop, Jonathan D; Davis, Raymond; Holz, Andres; Kolden, Crystal; Meddens, Arjan JH

Forest mastication treatments are increasingly utilized by land managers as a means of reducing tree cover for fire hazard mitigation and other habitat objectives in pinon-juniper (P-J) woodlands. Mastication converts trees into small pieces (e.g., wood chips), in the process creating canopy openings, redistributing fuel from the canopy to the surface, converting large diameter to small diameter fuels, and covering the ground with piles of woody debris. We measured vegetation and fuels at 192 sites in 24 pairs of 1–11-year-old mastication treatments and untreated adjacent controls in P-J woodlands of the Arkansas River valley, Colorado, and used paired t -tests, mixed-effects models, and gradient analysis (non-metric multidimensional scaling) to assess mastication effects. Treatments were associated with major, persistent ecological shifts relative to controls. Tree cover and canopy fuels were reduced in treatments; concomitantly, dead and down woody surface fuels, forb, and graminoid cover were elevated. Treatments exhibited much higher frequency, richness, and cover by a suite of non-native plant species including cheatgrass ( Bromus tectorum ). Non-native plant expansion appears linked to the disturbance associated with treatment activities, reductions in tree canopy, and alterations to ground cover, and effective mitigation of increases by these species may necessitate both pre- and post-treatment control measures. Shifts from native-dominated woodlands to open, weedy, herb- and shrub-dominated communities are likely to change patterns of abundance and habitat use by woodland- and forest-dependent wildlife. Decreased canopy fuels and increased herbaceous surface fuels including exotic annuals are expected to alter potential fire behavior. We encourage managers carrying out P-J mastication projects to explicitly address potential trade-offs between desired treatment outcomes and potentially unwelcome impacts, and how these might be mitigated. It may also be worth considering whether or not tree removal treatments will be warranted given anticipated climate change impacts to these woodlands.

Tree species are highly vulnerable to anthropogenic environmental change, and are increasingly being challenged by non-native pests and climate change. Rocky Mountain bristlecone pines are long-lived, exhibit delayed maturation, have low genetic diversity, and inhabit cold, high-elevation environments. They are threatened by the non-native disease white pine blister rust, warming temperatures, changing precipitation patterns, and altered disturbance regimes. The goal of this work was to (1) sample the range-wide genetic diversity of bristlecone pine for ex situ seed and tissue collections and research before the populations are impacted, (2) assess the health and ecological conditions of the species across its geographic range (61 stands) to provide a baseline by which to evaluate future changes, and (3) assess relationships between stand and regeneration characteristics and topographic, geographic, and climatic factors to identify vulnerabilities and proactive management interventions to increase population resilience. Local variation in topoclimate was strongly related to stand structure, composition, health, and regeneration. Bristlecone pine currently showed only minor impacts from insects and pathogens, but relationships between cone production and regeneration with climate variables suggest vulnerabilities to projected climate change. Both cone production and seedling regeneration were also linked to stand structure and ground cover that provide opportunities for proactive in situ management to increase population size to mitigate potential future climate- and pathogen-driven declines. These efforts represent the first proactive coordinated range-wide genetic collection design and forest health assessment for a threatened, but not yet impacted, tree species and offers a model for other species at risk.

Wildfires in forest ecosystems produce landscape mosaics that include relatively unaffected areas, termed fire refugia. These patches of persistent forest cover can support fire-sensitive species and the biotic legacies important for post-fire forest recovery, yet little is known about their abundance and distribution within fire perimeters. Readily accessible 30-m resolution satellite imagery and derived burn severity products are commonly employed to characterize post-fire landscapes, however, coarse image resolution, generalized burn severity thresholds, and other limitations can constrain accurate representation of fire refugia. This study quantifies the abundance and pattern of fire refugia within 10 fires occurring in ponderosa pine and dry mixed-conifer forests between 2000 and 2003. We developed high-resolution maps of post-fire landscapes using semi-automated, object-based classification of 1-m aerial imagery, conducted imagery- and field-based accuracy assessments, and contrasted these with Landsat-derived burn severity metrics. Fire refugia area within burn perimeters ranged from 20% to 57%. Refugia proportion generally decreased with increasing Landsat-derived burn severity, but still accounted for 3&ndash, 12% of areas classified as high severity. Patch size ranged from 1-m2 isolated trees to nearly 8000 ha, and median patch size was 0.01 ha&mdash, substantially smaller than a 30-m Landsat pixel. Patch size was negatively related to burn severity, distance to fire refugia from open areas was positively related to burn severity. Finally, optimized thresholds of 30-m post-fire normalized burn ratio (NBR) and relative differenced normalized burn ratio (RdNBR) delineated fire refugia with an accuracy of 77% when validated against the 1-m resolution maps. Estimations of fire refugia abundance based on Landsat-derived burn severity metrics are unlikely to detect small, isolated fire refugia patches. Finer-resolution maps can improve understanding of the distribution of forest legacies and inform post-fire management activities including reforestation and treatments.

Altered fire regimes can drive major and enduring compositional shifts or losses of forest ecosystems. In western North America, ponderosa pine and dry mixed‐conifer forest types appear increasingly vulnerable to uncharacteristically extensive, high‐severity wildfire. However, unburned or only lightly impacted forest stands that persist within burn mosaics—termed fire refugia—may serve as tree seed sources and promote landscape recovery. We sampled tree regeneration along gradients of fire refugia proximity and density at 686 sites within the perimeters of 12 large wildfires that occurred between 2000 and 2005 in the interior western United States. We used generalized linear mixed‐effects models to elucidate statistical relationships between tree regeneration and refugia pattern, including a new metric that incorporates patch proximity and proportional abundance. These relationships were then used to develop a spatially explicit landscape simulation model. We found that regeneration by ponderosa pine and obligate‐seeding mixed‐conifer tree species assemblages was strongly and positively predicted by refugia proximity and density. Simulation models revealed that for any given proportion of the landscape occupied by refugia, small patches produced greater landscape recovery than large patches. These results highlight the disproportionate importance of small, isolated islands of surviving trees, which may not be detectable with coarse‐scale satellite imagery. Findings also illustrate the interplay between patch‐scale resistance and landscape‐scale resilience: Disturbance‐resistant settings (fire refugia) can entrain resilience (forest regeneration) across the burn matrix. Implications and applications for land managers and conservation practitioners include strategies for the promotion and maintenance of fire refugia as components of resilient forest landscapes.

Natural resource managers are increasingly applying tree reduction treatments to piñon–juniper woodlands to meet a range of ecological, social, and economic goals. However, treatment effects on woodland-obligate bird species are not well understood. We measured multiscale avian occupancy on 29 paired (control/treatment) sites in piñon–juniper woodlands in central Colorado, USA. We conducted point counts at 232 stations, 3 times each season in 2014 and 2015. We used hierarchical multiscale modeling to obtain unbiased estimates of landscape and local occupancy (i.e. probability of use) in treated and untreated sites for 31 species. Treatments reduced the occupancy of conifer obligates, including Mountain Chickadee (Poecile gambeli), Clark’s Nutcracker (Nucifraga columbiana), and White-breasted Nuthatch (Sitta carolinensis), and increased occupancy of Lark Sparrow (Chondestes grammacus) and Mountain Bluebird (Sialia currucoides). Occupancy of Virginia’s Warbler (Oreothylpis virginiae) and Gray Flycatcher (Empidonax wrightii), two piñon–juniper specialists, decreased at the landscape scale in treated sites, and Pinyon Jay (Gymnorhinus cyanocephalus) occupancy decreased at the local scale. Tree reduction treatments in piñon–juniper woodlands have the potential to reduce habitat quality for a suite of bird species of conservation concern. We suggest that treatments designed to retain higher tree density and basal area will benefit conifer-obligate and piñon–juniper specialist bird species.

Extensive high‐severity wildfires have driven major losses of ponderosa pine and mixed‐conifer forests in the southwestern United States, in some settings catalyzing enduring conversions to non‐forested vegetation types. Management interventions to reduce the probability of stand‐replacing wildfire have included mechanical fuel treatments, prescribed fire, and wildfire managed for resource benefit. In 2011, the Las Conchas fire in northern New Mexico burned forested areas not exposed to fire for >100 yr, but also reburned numerous prescribed fire units and/or areas previously burned by wildfire. At some sites, the combination of recent prescribed fire and wildfire approximated known pre‐settlement fire frequency, with two or three exposures to fire between 1977 and 2007. We analyzed gridded remotely sensed burn severity data (differenced normalized burn ratio), pre‐ and post‐fire field vegetation samples, and pre‐ and post‐fire measures of surface fuels to assess relationships and interactions between prescribed fire, prior wildfire, fuels, subsequent burn severity, and patterns of post‐fire forest retention vs. conversion to non‐forest. We found that Las Conchas burn severity was lowest, and tree survival was highest, in sites that had experienced both prescribed fire and prior wildfire. Sites that had experienced only prescribed or prior wildfire exhibited moderate burn severity and intermediate levels of forest retention. Sites lacking any recent prior fire burned at the highest severity and were overwhelmingly converted to non‐forested vegetation including grassland, oak scrub, and weedy, herbaceous‐dominated types. Burn severity in the Las Conchas fire was closely linked to surface woody fuel loads, which were reduced by prior wildfire and prescribed fire. Our results support the restoration of fire regimes via prescribed fire and resource benefit wildfire to promote the resiliency of forest types vulnerable to fire‐mediated type conversion. The application of prescribed fire to reduce surface fuels following wildfire may reduce forest loss during subsequent fire under more extreme conditions. These findings are especially relevant given likely increases in vulnerability associated with climate change impacts to wildfire and forest dynamics.

Like many other high elevation alpine tree species, Rocky Mountain bristlecone pine (Pinus aristata Engelm.) may be particularly vulnerable to climate change. To evaluate its potential vulnerability to shifts in climate, we defined the suitable climate space for each of four genetic lineages of bristlecone pine and for other subalpine tree species in close proximity to bristlecone pine forests. Measuring changes in the suitable climate space for lineage groups is an important step beyond models that assume species are genetically homogenous. The suitable climate space for bristlecone pine in the year 2090 is projected to decline by 74% and the proportional distribution of suitable climate space for genetic lineages shifts toward those associated with warmer and wetter conditions. The 2090 climate space for bristlecone pine exhibits a bimodal distribution along an elevation gradient, presumably due to the persistence of the climate space in the Southern Rocky Mountains and exclusion at mid-elevations by conditions that favor the climate space of other species. These shifts have implications for changes in fire regimes, vulnerability to pest and pathogens, and altered carbon dynamics across the southern Rockies, which may reduce the likelihood of bristlecone pine trees achieving exceptional longevity in the future. The persistence and expansion of climate space for southern bristlecone pine genetic lineage groups in 2090 suggests that these sources may be the least vulnerable in the future. While these lineages may be more likely to persist and therefore present opportunities for proactive management (e.g., assisted migration) to maintain subalpine forest ecosystem services in a warmer world, our findings also imply heighted conservation concern for vulnerable northern lineages facing range contractions.

The spatial patterns resulting from large fires include refugial habitats that support surviving legacies and promote ecosystem recovery. To better understand the diverse ecological functions of refugia on burn mosaics, we used remotely sensed data to quantify neighborhood patterns of areas relatively unchanged following the 2011 Las Conchas fire. Spatial patterns of refugia measured within 10-ha moving windows varied across a gradient from areas of high density, clustered in space, to sparsely populated neighborhoods that occurred in the background matrix. The scaling of these patterns was related to the underlying structure of topography measured by slope, aspect and potential soil wetness, and spatially varying climate. Using a nonmetric multidimensional scaling analysis of species cover data collected post-Las Conchas, we found that trees and forest associates were present across the refugial gradient, but communities also exhibited a range of species compositions and potential functions. Spatial patterns of refugia quantified for three previous burns (La Mesa 1977, Dome 1996, Cerro Grande 2000) were dynamic between fire events, but most refugia persisted through at least two fires. Efforts to maintain burn heterogeneity and its ecological functions can begin with identifying where refugia are likely to occur, using terrain-based microclimate models, burn severity models and available field data.

The increasing frequency and severity of wildfires in semi-arid conifer forests as a result of global change pressures has raised concern over potential impacts on biodiversity. Ground-dwelling arthropod communities represent a substantial portion of diversity in conifer forests, and could be particularly impacted by wildfires. In addition to direct mortality, wildfires can affect ground-dwelling arthropods by altering understory characteristics and associated deterministic community assembly processes (e.g., environmental sorting). Alternatively, disturbances have been reported to increase the importance of stochastic community assembly processes (e.g., probabilistic dispersal and colonization rates). Utilizing pitfall traps to capture ground-dwelling arthropods within forest stands that were burned by one or two wildfires since 1996 in the Jemez Mountains of northern New Mexico, United States (USA), we examined the potential influences of deterministic versus stochastic processes on the assembly of these diverse understory communities. Based on family-level and genera-level arthropod identifications, we found that the multivariate community structures differed among the four fire groups surveyed, and were significantly influenced by the quantities of duff, litter, and coarse woody debris, in addition to tree basal area and graminoid cover. Taxon diversity was positively related to duff quantities, while taxon turnover was positively linked to exposed-rock cover and the number of logs on the ground. Despite the significant effects of these understory properties on the arthropod community structure, a combination of null modeling and metacommunity analysis revealed that both deterministic and stochastic processes shape the ground-dwelling arthropod communities in this system. However, the relative influence of these processes as a function of time since the wildfires or the number of recent wildfires was not generalizable across the fire groups. Given that different assembly processes shaped arthropod communities among locations that had experienced similar disturbances over time, increased efforts to understand the processes governing arthropod community assembly following disturbance is required in this wildfire-prone landscape.

Large and severe wildfires have raised concerns about the future of forested landscapes in the southwestern United States, especially under repeated burning. In 2011, under extreme weather and drought conditions, the Las Conchas fire burned over several previous burns as well as forests not recently exposed to fire. Our purpose was to examine the influences of prior wildfires on plant community composition and structure, subsequent burn severity, and vegetation response. To assess these relationships, we used satellite-derived measures of burn severity and a nonmetric multidimensional scaling of pre- and post- Las Conchas field samples. Earlier burns were associated with shifts from forested sites to open savannas and meadows, oak scrub, and ruderal communities. These non-forested vegetation types exhibited both resistance to subsequent fire, measured by reduced burn severity, and resilience to reburning, measured by vegetation recovery relative to forests not exposed to recent prior fire. Previous shifts toward non-forested states were strongly reinforced by reburning. Ongoing losses of forests and their ecological values confirm the need for restoration interventions. However, given future wildfire and climate projections, there may also be opportunities presented by transformations toward fire-resistant and resilient vegetation types within portions of the landscape.